Plant Nematodes in South Africa. 7. a Check List of Plant Nematodes from the Fynbos Biome, with a Description of Helicotylenchus Curatus Sp

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Plant nematodes in South Africa. 7. A check list of plant nematodes from the Fynbos Biome, with a description of Helicotylenchus curatus sp. n.

M. MARAIS, E. VAN DEN BERG, A. SWART and L.R. TIEDT

Marais, M., E. van den Berg, A. Swart and L.R. Tiedt. 2004. Plant nematodes in South Africa. 7. A check list of plant nematodes from the Fynbos Biome, with a description of Helicotylenchus curatus sp. n. Koedoe 47(1): 67–78. Pretoria. ISSN 0075-6458. Plant nematodes recorded during surveys in the Fynbos Biome are listed and a new Helicotylenchus species is described. Helicotylenchus curatus sp. n. is characterised by stylet length (42-46 µm in females, 37-40 µm in males), presence of two rudimentary subdorsal and two rudimentary subventral lobes on the labial disc, first lip annulus divided into six sectors, presence of fasciculi and presence of males. Nine families represented by 32 genera and 152 species were identified from the Fynbos Biome. The genera Criconema, Helicotylenchus, Hemicycliophora, Rotylenchus, Scutellonema and Xiphinema were found in more than 30 % of the localities, whereas Caloosia, Criconemoides, Ditylenchus, Geocenamus, Hemicriconemoides, Heterodera, Hoplo- laimus, Longidorus, Meloidogyne, Mesocriconema, Ogma, Paralongidorus, Paratri- chodorus, Paratylenchus, Pratylenchoides, Pratylenchus, Rotylenchulus, Trichodorus and Tylenchorhynchus were found at fewer localities. The genera Anguina, Hirschmaniella, Histotylenchus and Zygotylenchus were each identified from a single locality. Key words: biodiversity, Fynbos, Helicotylenchus, plant nematodes, new species, South Africa, taxonomy.

M. Marais, E. van den Berg, A. Swart, Nematology Unit, Biosystematics Division, ARC- Plant Protection Research Institute, Private Bag X134, Queenswood, 0121 Republic of South Africa; L.R. Tiedt, Laboratory for Electron Microscopy, North West University, Potchefstroom Campus, Potchefstroom, 2520 Republic of South Africa.

Introduction 2000) during the past 30 years. The Fynbos Biome is one of the biomes defined by Low In 1987, the South African Plant-Parasitic & Rebelo (1996). In South Africa, vegetation Nematode Survey (SAPPNS) programme structure and climate mostly define these was initiated with the aim to make a compre- ecological zones. The Fynbos Biome is con- hensive assessment of the nematode biodi- sidered by many to be synonymous with the versity resources of South Africa. One of the Cape Floral Kingdom, but this biome refers objectives of the SAPPNS is to compile an only to the two vegetation groups, viz., Fyn- inventory of the plant-parasitic nematodes of bos and Renosterveld within the region (Low South Africa. Although nematodes constitute & Rebelo 1996). Renosterveld is charac- an abundant and highly diverse group of terised by the dominance of members of the invertebrates, very little is known about their Asteraceae, specifically one species diversity in fynbos. The checklist reflects on Elytropappus rhinocerotis (renosterbos), collecting done by the Agricultural Research from which the vegetation type gets its name. Council, Western Cape Nature Conservation Although renosterbos is the characteristic Board, Rand Afrikaans University and Uni- dominant plant, other plants are also promi- versity of Stellenbosch (Heyns 1971; Kleyn- nent. These plants are all shrubs, charac- hans et al. 1996; Marais & Swart 1999; Van terised by small, tough grey leaves (Rebelo den Berg et al. 2003; Van den Berg & Tiedt 1996a). The various Fynbos vegetation types

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comprise most of the area of the Fynbos Results and discussion Biome. Fynbos is characterised by the pres- The classification of nematodes followed is ence of three elements, viz., a component based on Maggenti et al. (1988) for belonging to the Cape Reed family Tylenchina, Hunt (1993) for Aphelenchina (Restionaceae), an ericoid or heath compo- and Longidoridae and Decraemer (1995) for nent and a proteoid component. Fynbos is Trichodoriae. Nine families represented by also characterised by the presence of a num- 32 genera and 152 species were identified ber of endemic or near-endemic plant fami- from the Fynbos biome (Table 1). The gen- lies, viz., Bruniaceae (Blacktips), Geissolo- era Criconema, Helicotylenchus, Scutellone- maceae (Guyalone), Grubbiaceae (Sillyber- ma, Rotylenchus and Xiphinema were the ry), Penaeaceae (Brickleaf), Retziaceae most common and were found at 40 %, (Buttbush), Roridulaceae (Dewstick) and 35 %, 31 %, 30 % and 30 % of the localities, Stilbaceae (Candlestick) (Rebelo 1996b). respectively. The predominance of records in Helicotylenchus is a cosmopolitan genus the Mountain Fynbos vegetation type is a consequence of the number of samples col- with more than 200 species described lected in this vegetation type (Table 1). (Marais 2001). This genus is found in all the Mountain Fynbos is also the vegetation type biomes of South Africa. Thirty-one species with the highest total area (27 461 km2) of of Helicotylenchus have been reported from the ten Fynbos vegetation types (Low & South Africa. This paper is the result of an Rebelo 1996). ongoing investigation on the genus Helicoty- lenchus (Marais 1993, 1998, 2001; Marais & Buckley 1992; Marais & Quénéhervé 1996; Taxonomy Marais & Quénéhervé 1999; Marais et al. 2000; Van den Berg & Marais 1995; Van den Helicotylenchus curatus sp. n. Berg et al. 2003). Figs. 1 & 2 Measurements Material and methods Holotype female: L = 933 µm; a = 26.4; c = Soil samples were collected with a garden trowel, 43.8; c' = 1.2; stylet length = 42 µm; m = soil auger or spade to a depth of about 20-25 cm at 53 %; V = 62 %. each locality. Samples were collected in unprotected areas such as farms and plantations but also in nature Paratype females (n = 9): L = 1040 ± 110.1 reserves, wilderness areas and national parks. Nema- (839-1262) µm; a = 23.0 ± 2.9 (17.5–27.5); todes were extracted from 250 cm3 of soil, killed in c = 44.1 ± 4.5 (38.1–52.0); c' = 1.0 ± 0.2 water, preserved in FAA, TAF or FPG and mounted (0.8–1.2); stylet length = 44 ± 1.7 (42–46); in anhydrous glycerine (Hooper & Evans 1993; m = 54 ± 1.4 (52–56) %; V = 61 ± 1.8 Jenkins 1964; Kleynhans 1997; Koen & Furstenberg (58–64) %. 1970; Netscher & Seinhorst 1969; Seinhorst 1959). To obtain females of Meloidogyne for identification, Paratype males (n = 5): L = 864 ± 69.6 subsamples of each soil sample, in which root-knot (760–944) µm; a = 36.2 ± 3.4 (32.5–39.2); c nematode juveniles were found, were planted to = 30.0 ± 2.9 (27.0–34.5); c' = 1.8 ± 0.4 (1.4– tomato seedlings (cv. UC82B or Roma VF) in a 2.6); stylet length = 39 ± 1.4 (37–40) µm; greenhouse. After six weeks, females were extracted m = 57 ± 3.5 (52–60) %; spicule length = 33 from the roots and mounted in anhydrous glycerine ± 0.9 (32–34) µm; gubernaculum length = 10 (Kleynhans 1991). For scanning electron ± 0.7 (9–11) µm. microscopy, FPG-preserved specimens were used after dehydration in increasing concentrations of Female: Habitus spiral. Lip region 7 ± 0.5 acetone. Following conventional critical-point dry- (6–8) µm high and 10 ± 0.9 (10–13) µm ing and gold/palladium-coating (15 nm), specimens wide, truncate with distinct labial disc, six to were viewed with a Quanta 200 ESEM. seven annuli, basal annuli bulging out.

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Table 1 Plant nematodes found associated with the Fynbos Biome 1a 2 3 4 5 6 7 8 % Incidenceb Number of localities samples 1 12 16 28 159 2 1 7 ANGUINIDAE Anguina agrostis (Steinbuch, 1799) Filip'ev, 1936 + <1 Ditylenchus spp. + <1 Ditylenchus africanus Wendt, Swart, Vrain & Webster, 1995 + <1 BELONOLAIMIDAE Tylenchorhynchus spp. + + + + + 15 Tylenchorhynchus avaricus (Kleynhans, 1975) Fortuner & Luc, 1987 + <1 Tylenchorhynchus capitatus Allen, 1955 + + 1 Tylenchorhynchus clarus Allen, 1955 + <1 Tylenchorhynchus dewaelei Kleynhans, 1992 + <1 Tylenchorhynchus estherae Kleynhans, 1992 + + 1 Tylenchorhynchus indicus (Siddiqi, 1960) Fortuner & Luc, 1987 + <1 Tylenchorhynchus mashoodi Siddiqi & Basir, 1959 + <1 Tylenchorhynchus namibiensis Rashid & Heyns, 1990 + <1 Tylenchorhynchus phaseoli Sethi & Swarup, 1968 + <1 Histotylenchus spp. + <1 Geocenamus brevidens (Allen, 1955) Brzeski, 1991 + + 1 PRATYLENCHIDAE Hirschmaniella spp. + <1 Pratylenchoides spp. + + <1 Pratylenchus spp. + + + + + 11 Pratylenchus crenatus Loof, 1960 + <1 Pratylenchus delattrei Luc, 1958 + <1 Pratylenchus neglectus (Rensch, 1924) Filip'ev & Schuurmans Stekhoven, 1941 + + 1 Pratylenchus penetrans (Cobb, 1917) Filip'ev & Schuurmans Stekhoven, 1941 + + 3 Pratylenchus pratensis (de Man, 1880) Filip'ev, 1936 + <1 Pratylenchus vulnus Allen & Jensen, 1951 + <1 Pratylenchus zeae Graham, 1951 + 1 Zygotylenchus guevarai (Tobar Jiménez, 1963) Braun & Loof, 1966 + <1 HOPLOLAIMIDAE Hoplolaimus spp. + 3 Hoplolaimus capensis Van den Berg & Heyns, 1970 + + + 4 Hoplolaimus pararobustus (Schuurmans Stekhoven & Teunissen, 1938) Sher, 1963 + 1 Rotylenchus spp. + + 5 Rotylenchus alius Van den Berg, 1986c +++ 6 Rotylenchus brevicaudatus Colbran, 1962 + + + 8 Rotylenchus catharinae Van den Berg & Heyns, 1974 + + 1 Rotylenchus caudaphasmidius Sher, 1965 + 2 Rotylenchus incultus Sher, 1965 + + 3 Rotylenchus karooensis Van den Berg, 1986 + 1 Rotylenchus kenti Van den Berg, 1989 + + + 2 Scutellonema minutus (Sher, 1964) Germani, Baldwin & Wu, 1986 + <1

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Table 1 (continued) 1a 2 3 4 5 6 7 8 % Incidenceb Number of localities samples 1 12 16 28 159 2 1 7 Rotylenchus unisexus Sher, 1965 + + + 6 Rotylenchus usitatus Van den Berg & Heyns, 1974 + <1 Helicotylenchus spp. + + + + + 22 Helicotylenchus californicus Sher, 1966 + + 2 Helicotylenchus cavenessi Sher, 1966 + <1 Helicotylenchus curatus sp. n. + <1 Helicotylenchus digonicus Perry in Perry, Darling & Thorne, 1959 + + 2 Helicotylenchus dihystera (Cobb, 1893) Sher, 1961 + + + 4 Helicotylenchus exallus Sher, 1966 + + 6 Helicotylenchus hydrophilus Sher, 1966 + <1 Helicotylenchus minzi Sher, 1966 + 4 Helicotylenchus paraplatyurus Siddiqi, 1972 + + 1 Helicotylenchus pseudorobustus (Steiner, 1914) Golden, 1956 + 1 Helicotylenchus serenus Siddiqi, 1963 + + 1 Scutellonema spp. + + + + 8 Scutellonema bizanae Van den Berg & Heyns, 1973 + + + + 13 Scutellonema brachyurus (Steiner, 1938) Andrássy, 1958 + + + + 9 Scutellonema nigermontanum Van den Berg, 1990 + <1 Scutellonema tsitsikamense Van den Berg, 1976 + 1 Scutellonema unum Sher, 1964 + <1 Rotylenchulus spp. + 1 Rotylenchulus borealis Loof & Oostenbrink, 1962 + 2 Rotylenchulus macrodoratus Dasgupta, Raski & Sher, 1968 + 1 Rotylenchulus parvus (Williams, 1960) Sher, 1961 + + + + 5 Rotylenchulus reniformis Linford & Oliveira, 1940 + HETERODERIDAE Heterodera spp. + 1 Heterodera trifolii Goffart, 1932 + <1 Meloidogyne spp. + + + 5 Meloidogyne hapla Chitwood, 1949 + + 3 Meloidogyne incognita (Kofoid & White, 1919) Chitwood, 1949 + 3 Meloidogyne javanica (Treub, 1885) Chitwood, 1949 + + + 3 CRICONEMATIDAE Criconema spp. + + + + 13 Criconema ananas (Heyns, 1970) Siddiqi, 1986 + <1 Criconema corbetti (De Grisse, 1967) Raski & Luc, 1985 + 3 Criconema crassianulatum (de Guiran, 1963) Raski & Luc, 1985) + 1 Criconema duplicivestitum (Andrássy, 1963) Raski & Luc, 1985 + 11 Criconema indigenae Van den Berg & Meyer, 1991c +1 Criconema mutabile (Taylor, 1936) Raski & Luc, 1987 + + + + + 4 Criconema proteae Van den Berg & Meyer, 1991c +1 Criconema sanctifrancisci (Van den Berg & Heyns, 1977) Raski & Luc, 1985 + + 11 Criconema simplex Marais & Van den Berg, 1996c +1

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Table 1 (continued) 1a 2 3 4 5 6 7 8 % Incidenceb Number of localities samples 1 12 16 28 159 2 1 7 Criconema sirgeli Van den Berg & Meyer, 1987c +<1 Criconemoides parvus Raski, 1952 + + 2 Hemicriconemoides spp. + <1 Hemicriconemoides brachyurus (Loos, 1949) Chitwood & Birchfield, 1957 + + + + 6 Hemicriconemoides capensis Van den Berg, 1990 + <1 Hemicriconemoides cedrusmontanus Van den Berg & Meyer, 1991c +<1 Mesocriconema spp. + + + + + + 12 Mesocriconema ferniae (Luc, 1959) Loof & De Grisse, 1989 + 3 Mesocriconema obtusicaudatum (Heyns, 1962) Loof & De Grisse, 1989 + + + 3 Mesocriconema sphaerocephalum (Taylor, 1936) Loof & De Grisse, 1989 + 1 Mesocriconema thabaum Van den Berg, 1996c +<1 Mesocriconema xenoplax (Raski, 1952) Loof & De Grisse, 1989 + + + 1 Ogma spp. + <1 Ogma civellae civellae Reay & Davies, 1998 + + Ogma decalineatum (Chitwood, 1957) Andrássy, 1979 + 1 Ogma inornatum (Van den Berg, 1983) Siddiqi, 1986c +<1 Ogma rhombosquamatum (Mehta & Raski, 1971) Andrássy, 1979 + + 1 Ogma squamiferum (Heyns, 1970) Andrássy, 1979c ++ 1 Caloosia exigua Van den Berg, Marais & Tiedt, 2003c + Caloosia peculiaris Van den Berg & Meyer, 1991c +<1 Hemicycliophora spp. + + + 9 Hemicycliophora demani Edward & Rai, 1971 + 2 Hemicycliophora epicharoides Loof, 1968 + <1 Hemicycliophora halophila Yeates, 1967 + + 1 Hemicycliophora labiata Colbran, 1960 + + + 7 Hemicycliophora natalensis Loof & Heyns, 1969 + + 1 Hemicycliophora nullinca Van den Berg, 1987 + <1 Hemicycliophora peca Van den Berg, 1987c +<1 Hemicycliophora stiaani Van den Berg & Tiedt, 1999c +<1 Hemicycliophora typica de Man, 1921 + 1 Hemicycliophora wesca Van den Berg & Meyer, 1987c +<1 TYLENCHULIDAE Paratylenchus spp. + + <1 Paratylenchus arculatus Luc & de Guiran, 1962 + <1 Paratylenchus elachistus Steiner, 1949 + <1 Paratylenchus projectus Jenkins, 1956 + + 1 Paratylenchus vandenbrandei De Grisse, 1962 + <1 TRICHODORIDAE Trichodorus spp. + 5 Trichodorus iuventus Decraemer & Marais, 2000c +<1

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Table 1 (continued) 1a 2 3 4 5 6 7 8 % Incidenceb Number of localities samples 1 12 16 28 159 2 1 7 Trichodorus philipi De Waele & Van Mieghem, 1990c +1 Trichodorus vandenbergae De Waele & Kilian, 1992 + 2 Paratrichodorus spp. + + 4 Paratrichodorus lobatus (Colbran. 1956) Siddiqi, 1974 + + + 3 Paratrichodorus minor (Colbran. 1956) Siddiqi, 1974 + + + 3 Paratrichodorus porosus (Allen, 1957) Siddiqi, 1974 + <1 LONGIDORIDAE Paralongidorus spp. + 4 Paralongidorus capensis (Heyns, 1967) Liebenberg, Heyns & Swart, 1993 + <1 Paralongidorus christiani Liebenberg, Heyns & Swart, 1993c +1 Paralongidorus costatus (Jacobs & Heyns, 1987) Siddiqi, Baujard & Mounport, 1993 + <1 Paralongidorus deborae (Jacobs & Heyns, 1982) Luc & Doucet, 1984 + <1 Paralongidorus spasskii Heyns, 1972 + 1 Longidorus spp. + + 2 Longidorus fursti Heyns, Coomans, Hutsebaut & Swart, 1987 + <1 Longidorus jagerae Heyns & Swart, 1998 + <1 Longidorus juvenilis Dalmasso, 1969 + + 1 Longidorus pisi Edward, Misra & Singh, 1964 6 Xiphinema spp. + + + + 8 Xiphinema americanum Cobb, 1913 + 1 Xiphinema barbercheckae Coomans & Heyns, 1985 + + + + 3 Xiphinema bolandium Coomans & Heyns, 1985c +++ 5 Xiphinema capense Coomans & Heyns, 1985 + 2 Xiphinema diffusum Lamberti & Bleve-Zacheo, 1979 + + 1 Xiphinema elongatum Schuurmans Stekhoven & Teunissen, 1938 + + + 1 Xiphinema hardingi Joubert, Kruger & Heyns, 1988 + 1 Xiphinema krugi Lordello, 1955 + + 1 Xiphinema mampara Heyns, 1979 + <1 Xiphinema meridianum Heyns, 1971 + + + 4 Xiphinema mluci Heyns, 1976 + <1 Xiphinema pachtaicum (Tulaganov, 1938) Kir'yanova, 1951 + + 2 Xiphinema parvistilus Heyns, 1971 + + 2 Xiphinema ripogranum Hutsebaut, Heyns & Coomans, 1988c ++ + 6 Xiphinema vanderlindei Heyns, 1962 + + + 1 Xiphinema variabile Heyns, 1966 + <1 Xiphinema vitis Heyns, 1974 + + 1 Xiphinema zulu Heyns, 1965 + <1 a Vegetation types of the Fynbos Biome according to Low & Rebelo (1996) (1 = Escarpment Mountain Renos- terveld, 2 = Central Mountain Renosterveld, 3 = West Coast Renosterveld, 4 = South and South-west Coast Renosterveld, 5 = Mountain Fynbos, 6 = Grassy Fynbos, 7 = Laterite Fynbos, 9 = Sand Plain Fynbos) b Percentage incidence in total number of samples collected during surveys. c Nematodes described from the Fynbos Biome.

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a b

c d

Fig. 1. Helictylenchus curatus sp. n. Female. (a) Habitus. (b) En face view, lip region. (c) Lateral view, anterior part of body. (d) Lateral field. (e-f) Posterior part of body. (Scale bar: a = 100 µm, b-e = 4 µm).

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Labial disc rectangular in en face view, with Male: Habitus C-shaped (20 %) to spiral two rudimentary subdorsal and two rudimen- (80 %). Lip region 6 ± 0.5 (5–6) µm high and tary subventral lobes. First lip annulus divid- 8 ± 0.4 (8–9) µm wide; with six to seven ed into six sectors, two lateral, two subven- annuli, basal annuli bulging out. Anterior tral and two subdorsal. Outer margins of the cephalids three to four annuli posterior to well-developed labial framework extend 4 ± basal ring, posterior cephalids 12 to 14 0.2 (3–4) µm backward from basal plate. annuli posterior to anterior cephalids. Stylet Anterior cephalids three to four annuli poste- more slender than that of female; stylet rior to basal plate, posterior cephalids 11 to knobs 2 ± 0.4 (1–2) µm high and 4 ± 0.8 13 annuli posterior to anterior cephalids. (4–5) µm wide, anterior faces flattened Stylet slender, stylet knobs 4 ± 0.5 (3–4) µm (50 %) or indented (50 %). Position of DGO high and 7 ± 1.3 (6–10) µm wide; anterior 4 ± 0.6 (3–4) µm behind stylet knobs. Medi- faces rounded (14 %), flattened (29 %), an bulb oval, 15 ± 2.3 (13–19) µm long and indented (14 %) or flattened and slightly 10 µm wide; valve 3 ± 0.3 (3–4) µm long inclined backward (43 %). Position of dorsal and 3 ± 0.8 (2–4) µm wide. Length of gland opening (DGO) 5 ± 1.7 (3–9) µm oesophagus 146 ± 9.3 (133–153) µm, with behind stylet knobs. Median bulb oval, 16 length to end of glands 169 ± 15.4 (147– ± 1.4 (14–18) µm long and 12 ± 1.7 184) µm. Oesophagus with 29 ± 8.9 (18– (10–15) µm wide; valve 4 ± 0.4 (3–4) µm 39) µm long ventral or ventro-lateral over- long and 3 µm wide. Length of oesophagus lap. Excretory pore 129 ± 5.4 (121–136) µm 151 ± 13.0 (134–175) µm, with length to end from front, i.e. at 15 ± 0.7 (14–16) % of body of gland 172 ± 19.0 (154–212) µm. Oesoph- length. Hemizonid one to three annuli long, agus with 22 ± 4.3 (16–28) µm long ventral located two to three annuli anterior to excre- and dorsal overlap. Excretory pore 154 ± tory pore. Hemizonion half to one annuli 11.0 (142–172) µm from front, i.e. at 15 ± long, located 10 to 13 annuli posterior to 1.3 (14–17) % of body length. Hemizonid excretory pore. Fasciculi present. Width of two annuli long located one to three annuli annuli at midbody 2 µm. Body width at anterior to excretory pore. Hemizonion half excretory pore 20 ± 1.8 (18–23) µm, at mid- an annulus long, located 10 annuli posterior body 24 ± 3.3 (21–29) µm and at cloaca 17 to excretory pore (n = 1). Fasciculi present. ± 3.3 (15–24) µm. Lateral field 6 ± 0.3 Width of annulus at midbody 2 µm. Body (6–7) µm wide; outer two lines areolated width at excretory pore 33 ± 4.7 (26–40) µm, anterior to and opposite bursa, crenate or at midbody 46 ± 8.2 (35–59) µm and at anus incompletely areolated on body. Phasmids 24 ± 4.8 (18–29) µm. Two branches of the located posterior to cloaca. Margin of bursa reproductive system both functional, length almost straight, flattened. Tail 31 ± 4.9 of posterior branch 81 ± 16.6 (57–94) % of (26–39) µm long, conical with rounded pro- corresponding anterior branch length; anteri- jection. or branch 276 ± 59.8 (165–334) µm and pos- Diagnosis terior branch 199 ± 38.5 (156–235) µm long. Spermatheca set off, empty or filled with H. curatus sp. n. is characterised by stylet sperm. Epiptygma folded into vagina. Later- length (42–46 µm in females, 37–40 µm in al field 10 ± 1.7 (8–13) µm wide; outer two males), presence of two rudimentary subdor- lines areolated opposite oesophagus, crenate sal and two rudimentary subventral lobes on or incompletely areolated on rest of body; the labial disc, first lip annulus divided into inner two lines end on tail in a u-shaped six sectors, presence of fasciculi and pres- (80 %) or open m-shaped pattern (20 %). ence of males. Phasmids located from five annuli posterior Relationships to opposite anus. Tail 24 ± 2.3 (20–28) µm long, asymmetrical, more curved dorsally, A conspicuous character of H. curatus sp. n. with rounded end, with 13 to 18 ventral is the short DGO (5–9 µm) in females and annuli, some annuli irregular. (3–4 µm) in males. This short DGO fit into

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a b c

20 µm

f g h

Fig. 2. Helictylenchus curatus sp. n. Female. (a) Oesophageal region. (d) Lateral field and fasciculi. (e) Vul- val region. (f-h) Posterior part of body. Male. (b) Oesophageal region. (c) Posterior part of body.

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the diagnosis for the genus where the DGO continuous), labial disc characteristics (with varies from 3 µm (H. labiatus in Yeates & two rudimentary subdorsal and two rudimen- Wouts 1992) to 23 µm (H. erythrinae in tary subventral lobes present vs without any Marais 2001). According to Fortuner (1984), lobes), number of lip annuli (six to seven vs this character has little value for identifica- four to five), presence vs absence of fasci- tion, therefor the DGO was not used as a dif- ferentiating character. The stylet length has culi, a-value (17.5–27.5 vs 35–40), stylet the smallest coefficient of variation among length (42–46 µm vs 39–42 µm), m-value the quantitative characters (Fortuner 1979; (52–56 % vs 48–50 %), position of phasmids Fortuner et al. 1981). There is a group of (opposite anus to five annuli posterior to species in Helicotylenchus characterised by anus vs five to eight annuli anterior to anus) long bodies and stylets, the respective mea- and tail form (asymmetrical, more curved surements ranging from 800 µm to 1300 µm dorsally, with rounded end vs straight, taper- and from 35 µm to 46 µm. Nine species are ing dorsally with unstriated ventral portion). included in this group: H. arliani Khan, The males differ in body length (760– Singh & Lal, 1998, H. canalis Sher, 1966, H. coomansi Sharafati-Ali & Loof, 1975, 944 µm vs 1190–1260 µm), position of H. dolichodoryphorus Sher, 1966, H. kash- excretory pore from front (121–126 µm vs mirensis Fotedar & Handoo, 1974, 161 µm) and tail length (26–39 µm vs 41 µm H. macrostylus Marais & Quénéhervé, 1996, — calculated from paratype material.). The H. orthosomaticus Siddiqi, 1972, H. para- new species can be separated from H. canalis Sauer & Winoto, 1975, H. rohtangus macrostylus in presence of fasciculi vs Jairajpuri & Baqri, 1973 and H. tunisiensis absence, m-value (52–56 % vs 42–45 %), lip Siddiqi, 1963 (Jairajpuri & Baqri 1973; region shape (truncate with basal annuli Fotedar & Handoo 1974; Kepenekci & Ökten 1996; Khan et al. 1998; Marais 1998; bulging out vs continuous), labial disc char- Marais & Quénéhervé 1996; Marais et al. acteristics (rectangular, with two rudimenta- 2003; Sharafati-Ali & Loof 1975; Sher 1966; ry subdorsal and two rudimentary subventral Siddiqi 1972; Van den Berg & Kirby 1979). lobes vs round with no lobes), first lip annu- H. curatus sp. n. was compared with three of lus divided vs not divided. these species, where the stylet lengths are more than 40 µm i.e. H. canalis, H. cooman- Type locality and habitat si and H. macrostylus. The new species can Collected from Cyclopia plicata Kies at be separated from H. canalis in lip region Haarlem, Langkloof, Western Cape Province shape (truncate with basal annuli bulging out (33°44'S, 23°20'E) in a Mountain Fynbos vs flattened to rounded, not set off), labial disc characteristics (rectangular in en face vegetation type by H. Hugo. view, with two rudimentary subdorsal and Type material two rudimentary subventral lobes vs oval in en face view without any lobes), first lip Holotype female (slide number 36524), six annulus divided vs not divided, mean body female paratypes and seven male paratypes length (1040 µm vs 859 µm), position of (slide 36525–36528) deposited in the excretory pore from front (142–172 µm vs National Collection of Nematodes, Biosys- 121–143 µm), mean tail length (24 µm vs tematics Division, ARC-Plant Protection 20 µm), position of phasmids (opposite anus Research Institute, Pretoria, South Africa. to five annuli posterior to anus vs two to seven annuli anterior to anus) and presence Etymology of males vs absence of males. Females of the new species can be distinguished from From the Latin cura, meaning to care, in H. coomansi females in lip region shape recognition of Liezel Scheepers for her (truncate with basal annuli bulging out vs invaluable assistance.

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Acknowledgements from Nepal. International Journal of Nematol- ogy 8: 27–32. We thank N.H. Buckley and E. van Niekerk (ARC- KOEN, H. & J.P. FURSTENBERG. 1970. 'n Beknopte PPRI) for technical assistance, the various farmers, laboratoriumhandleiding vir nematologie. Pre- foresters and officers in charge for their help in toria: Departement van Landbou-Tegniese Dien- collecting the soil samples and P.A.A. Loof ste. (Tegniese Mededeling 92.) (Landbouwuniversiteit Wageningen) for the loan of KLEYNHANS, K.P.N. 1991. The root-knot nematodes H. coomansi paratypes. of South Africa. Pretoria: Department of Agri- cultural Development, South Africa. (Technical Communications no. 195.) References KLEYNHANS, K.P.N. 1997. Collecting and preserving nematodes. A manual for a SAFRINET course in practical nematology. ARC-Plant Protection DECRAEMER, W. 1995. The family Trichodoridae: Research Institute. Pretoria. Stubby root and virus vector nematodes. KLEYNHANS, K.P.N., E. VAN DEN BERG, A. SWART, M. Dorderecht: Kluwer. MARAIS & N.H. BUCKLEY. 1996. Plant nema- FORTUNER, R. 1979. Morphometrical variability in todes in South Africa. Pretoria: ARC-Plant Pro- Helicotylenchus Steiner, 1945. I. The progeny of tection Research Institute. (Plant Protection a single female. Revue de Nématologie 2: 179–202. Research Institute Handbook; no. 8.) LOW, A.B. & A.G. REBELO. 1996. Vegetation of South FORTUNER, R. 1984. Morphometrical variability in Helicotylenchus Steiner, 1945. 6: Value of the Africa, Lesotho and Swaziland. A companion to characters used for specific identification. Revue the Vegetation Map of South Africa, Lesotho and de Nématologie 7: 245–264. Swaziland. Pretoria: Department of Environ- mental Affairs & Tourism. FORTUNER, R., G. MERNY & C. ROUX. 1981. Mor- phometrical variability in Helicotylenchus Stein- MAGGENTI, A.R., M. LUC, D.J. RASKI, R. FORTUNER er, 1945. 3: Observations on African populations & E. GERAERT. 1988. A reappraisal of Tylenchi- of Helicotylenchus dihystera and considerations na (Nemata). 11. List of generic and supra- on related species. Revue de Nématologie 4: generic taxa, with their junior synonyms. Revue 235–260. de Nématologie 11: 177–188. FOTEDAR, D.N. & Z.A. HANDOO. 1974. Two new MARAIS, M. 1993. On some Helicotylenchus Steiner, species of Helicotylenchus (Hoplolaiminae: 1945 from South Africa. Phytophylactica 25: Nematoda) from Kashmir, India. Journal of Sci- 21–35. ence Kashmir University 2: 57–62. MARAIS, M. 1998. Some species of Helicotylenchus HEYNS, J. 1971. A guide to the plant and soil nema- Steiner, 1945 from South Africa (Nematoda: todes of South Africa. Cape Town: Balkema. Hoplolaimidae). Fundamental and Applied HOOPER, D.J. & K. EVANS. 1993. Extraction, identi- Nematology 21: 327–352. fication and control of plant nematodes. Pp. MARAIS, M. 2001. A monograph of the genus Heli- 1–59. In: EVANS, K., D.L. TRUDGILL & J.M. cotylenchus Steiner, 1945 (Nemata: Hoplolaimi- WEBSTER (eds.). Plant parasitic nematodes in dae). Ph.D. Agric. dissertation, University of temperate agriculture. Wallingford: CAB Inter- Stellenbosch, Stellenbosch. national. MARAIS, M. & N.H. BUCKLEY. 1992. External mor- HUNT, D.J. 1993. Aphelenchida, Longidoridae and phology of eight South African Helicotylenchus Trichodoridae: Their systematics and bionom- species (Hoplolaimidae: Nemata). Phytophylac- ics. Wallingford: CAB International. tica 24: 297–306. JAIRAJPURI, M.S. & Q.H. BAQRI. 1973. Nematodes of MARAIS, M. & P. QUÉNÉHERVÉ. 1996. Helicoty- high altitudes in India. I. Four new species of lenchus macrostylus n.sp. (Hoplolaimidae: Tylenchida. Nematologica 19: 19–30. Nemata) from French Guiana with notes on JENKINS, W.R. 1964. A rapid centrifugal-flotation three known species. Nematropica 26: 39–45. technique for separating nematodes from soil. MARAIS, M. & P. QUÉNÉHERVÉ. 1999. A new species Plant Disease Reporter 48: 692. of Helicotylenchus from French Guiana, with KEPENEKCI, I. & M.E. ÖKTEN. 1996. Beypazari notes on two known species (Nemata: Hoplo- (Ankara) Ilçesi'nde havuf ile münabeye giren laimidae). Journal for Nematode Morphology domates ekiliú alanlarinda saptanan Helicoty- and Systematics 2: 81–88. lenchus (Tylenchida, Hoplolaimidae) cinsine MARAIS, M., P. QUÉNÉHERVÉ, L.R. TIEDT & A.J. bagli türler. Türk Entomoloji Dergisi 20: MEYER. 2003. Description of Helicotylenchus 137–148. marethae n.sp., with redescriptions of some KHAN, E., M. SINGH & M. LAL. 1998. Four new Helicotylenchus species (Nemata: Hoplolaimi- species of tylenchids (Nematoda: Tylenchida) dae) from South Africa and Martinique. Journal

ISSN 0075-6458 77 Koedoe 47/1 (2004) marais.qxd 2004/04/15 07:57 Page 78

of Nematode Morphology and Systematics 5: toda: Hoplolaimidae). Nematologica 21: 1–19. (2002 published in 2003) 207–212. MARAIS, M. & A. SWART. 1999. Plant nematodes in SHER, S.A. 1966. Revision of the Hoplolaiminae the Lottering Plantation, Eastern Cape Province. (Nematoda) VI. Helicotylenchus Steiner, 1945. South African Forestry Journal 186: 77–79. Nematologica 12: 1–56. MARAIS, M. & E. VAN DEN BERG, P. QUÉNÉHERVÉ & SIDDIQI, M.R. 1972. On the genus Helicotylenchus L.R. TIEDT. 2000. Description of Helicoty- Steiner, 1945 (Nematoda: Tylenchida), with lenchus kermarreci n.sp., with notes on some descriptions of nine new species. Nematologica Helicotylenchus Steiner, 1945 and a Rotylenchus 18: 74–91. Filip'ev, 1936 species (Nemata: Hoplolaimidae) VAN DEN BERG, E. & M. F. KIRBY. 1979. Some spi- from the Guadeloupe Islands, French West Indies. Journal for Nematode Morphology and ral nematodes from the Fiji Islands (Hoplolaim- Systematics 2: 159–172. (1999 published in idae: Nematoda). Phytophylactica 11: 99–109. 2000). VAN DEN BERG, E. & M. MARAIS. 1995. New species NETSCHER, C. & J.W. SEINHORST. 1969. Propionic of Tylenchina Chitwood (Nemata) from the acid better than acetic acid for killing nema- Amazonas province, Brazil. African Plant Pro- todes. Nematologica 15: 286. tection 1: 25–39. REBELO, A.G. 1996a. Renosterveld. Pp. 63. In: LOW, VAN DEN BERG, E., M. MARAIS, S. GAIDASHOVA & A.B. & A.G. REBELO. Vegetation of South Africa, L.R. TIEDT. 2003. Hoplolaimidae Filip’ev, 1934 Lesotho and Swaziland. A companion to the Veg- (Nemata) from Rwandan banana fields. African etation Map of South Africa, Lesotho and Swazi- Plant Protection 9: 31–42. land. Pretoria: Department of Environmental VAN DEN BERG, E., M. MARAIS & L.R. TIEDT. 2003. Affairs & Tourism. Plant nematodes in South Africa. 5. Plant nema- REBELO, A.G. 1996b. Fynbos. Pp. 68–70. In: LOW, todes from the Gouekrans area of the Swartberg A.B. & A.G. REBELO. Vegetation of South Africa, nature Reserve, with descriptions of one new Lesotho and Swaziland. A companion to the Veg- and one known Hemicycliophorionae (Nemata). etation Map of South Africa, Lesotho and Swazi- African Plant Protection 9: 43–52. land. Pretoria: Department of Environmental VAN DEN BERG, E. & L.R. TIEDT. 2000. Hemicyclio- Affairs & Tourism. phora stiaani sp.n. with notes on three tylenchid SAUER, M. R. & R. WINOTO. 1975. The genus Heli- cotylenchus Steiner, 1945 in West Malaysia. (Nemata) species from South Africa. Journal of Nematologica 21: 341–350. Nematode Morphology and Systematics 2: SEINHORST, J.W. 1959. A rapid method for the trans- 137–148. (1999 published 2000). fer of nematodes from fixative to anhydrous YEATES, G.W. & W.M. WOUTS. 1992. Helicoty- glycerin. Nematologica 4: 67–69. lenchus spp. (Nematoda: Tylenchida) from man- SHARAFATI-ALI, S. & P.A.A. LOOF. 1975. Two new aged soils in New Zealand. New Zealand Jour- species of Helicotylenchus Steiner, 1945 (Nema- nal of Zoology 19: 13–23.

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